Nitrogen Dynamics in Ice Storm-Damaged Forest Ecosystems: Implications for Nitrogen Limitation Theory

Despite the widely recognized importance of disturbance in accelerating the loss of elements from land, there have been few empirical studies of the effects of natural disturbances on nitrogen (N) dynamics in forest ecosystems. We were provided the unusual opportunity for such study, partly because the intensively monitored watersheds at the Hubbard Brook Experimental Forest (HBEF), New Hampshire, experienced severe canopy damage following an ice storm. Here we report the effects of this disturbance on internal N cycling and loss for watershed 1 (W1) and watershed 6 (W6) at the HBEF and patterns of N loss from nine other severely damaged watersheds across the southern White Mountains. This approach allowed us to test one component of N limitation theory, which suggests that N losses accompanying natural disturbances can lead to the maintenance of N limitation in temperate zone forest ecosystems. Prior to the ice storm, fluxes of nitrate (NO₃ ^–) at the base of W1 and W6 were similar and were much lower than N inputs in atmospheric deposition. Following the ice storm, drainage water NO₃ ^– concentrations increased to levels that were seven to ten times greater than predisturbance values. We observed no significant differences in N mineralization, nitrification, or denitrification between damaged and undamaged areas in the HBEF watersheds, however. This result suggests that elevated NO₃ ^- concentrations were not necessarily due to accelerated rates of N cycling by soil microbes but likely resulted from decreased plant uptake of NO₃ ^-. At the regional scale, we observed high variability in the magnitude of NO₃ ^- losses: while six of the surveyed watersheds showed accelerated rates of NO₃ ^– loss, three did not. Moreover, in contrast to the strong linear relationship between NO₃ ^– loss and crown damage within HBEF watersheds [r ²: (W1 = 0.91, W6 = 0.85)], stream water NO₃ ^– concentrations were weakly related to crown damage (r ² = 0.17) across our regional sites. The efflux of NO₃ ^– associated with the ice storm was slightly higher than values reported for soil freezing and insect defoliation episodes, but was approximately two to ten times lower than NO₃ ^– fluxes associated with forest harvesting. Because over one half of the entire years worth of N deposition was lost following the ice storm, we conclude that catastrophic disturbances contribute synergistically to the maintenance of N limitation and widely observed delays of N saturation in northern, temperate zone forest ecosystems. Present address: Department of Ecology and Evolutionary Biology, Princeton University, Guyot Hall, Princeton, New Jersey 08544, USA.     

Characteristics of mineral particles in the human bone/cartilage interface

Bone and cartilage consist of different organic matrices, which can both be mineralized by the deposition of nano-sized calcium phosphate particles. We have studied these mineral particles in the mineralized cartilage layer between bone and different types of cartilage (bone/articular cartilage, bone/intervertebral disk, and bone/growth cartilage) of individuals aged 54 years, 12 years, and 6 months. Quantitative backscattered electron imaging and scanning small-angle X-ray scattering at a synchrotron radiation source were combined with light microscopy to determine calcium content, mineral particle size and alignment, and collagen orientation, respectively. Mineralized cartilage revealed a higher calcium content than the adjacent bone (p<0.05 for all samples), whereas the highest values were found in growth cartilage. Surprisingly, we found the mineral platelet width similar for bone and mineralized cartilage, with the exception of the growth cartilage sample. The most striking result, however, was the abrupt change of mineral particle orientation at the interface between the two tissues. While the particles were aligned perpendicular to the interface in cartilage, they were oriented parallel to it in bone, reflecting the morphology of the underlying organic matrices. The tight bonding of mineralized cartilage to bone suggests a mechanical role for the interface of the two elastically different tissues, bone and cartilage.     

Effects of delayed hay removal on the nutrient balance of roadside plant communities

1. Mass losses and nutrient losses from fresh roadside cuttings were studied in the field during a 6-week period. Large amounts (over 50%) appear to be lost from the cuttings. The losses were positively related to initial nutrient concentrations during this short-term study. Mass and nitrogen losses were best explained by the initial C:N ratio, phosphorus and potassium losses by the initial phosphorus concentration.
2. For potassium the losses were particularly large (up to 90%). For this element only, the observed relationship between loss rate and initial chemical composition could not be established significantly. It is concluded that potassium is mainly lost by leaching whereas the major nitrogen, phosphorus and mass losses are most probably caused by rapid microbial decomposition of readily soluble substances.
3. Using existing data on chemical composition of other roadside cuttings, nutrient losses after different hay removal delay times were modelled for different plant communities. All or most of the losses were assumed to return to the soil system.
4. When soil impoverishment is aimed for, cuttings should be removed within 1 or 2 weeks in most plant communities. If removal is delayed longer, the amounts of nutrients removed will often fall below the annual atmospheric input. In plant communities where annual above-ground production of nitrogen and phosphorus are lower than the annual atmospheric deposition already, rapid removal of the cuttings may be the only way to maintain at least potassium at a limiting level.
5. The main effect of hay-making on the soil nutrient status most likely consists of a reduction of the potassium availability, at least on sandy soils with a low cation exchange capacity and provided there is little delay in hay removal.     

Nitrogen-15 labeling effectiveness of two tropical legumes

Nitrogen-15 foliar applications for the production of field-labeled plant tissues may achieve more effective labeling of plant shoot and root tissues and minimize directly labeling the soil N fraction as occurs when¹⁵ N is soil applied. Consequently, foliar-labeled plant tissues should be better suited for subsequent ¹⁵N mineralization studies. A field experiment was conducted to determine the effectiveness of ¹⁵N-labeling and the accumulation of ¹⁵N in various plant parts of two tropical legumes. Desmodium ovalifolium Guillemin and Perrottet and Pueraria phaseoloides (Roxb.) Benth., grown in 0.5 m² microplots, were labeled with foliar-applied urea containing 99 atom% ¹⁵N. Plants in each microplot received a total of 0.1698 g ¹⁵N that was applied all at once or split equally into two, three or four applications. Legume shoots and roots and soil were destructively harvested and analyzed for total ¹⁵N content. Averaged over both legumes and foliar application rates, total plant (shoots, flowers, leaf litter, and roots) recovery was approximately 79% of the ¹⁵N applied. The soil contained 3% of the ¹⁵N applied, of which 2.5 and 0.5% were in the inorganic and organic fractions, respectively. Nitrogen-15 recovery in shoots (76%) was sixty-five fold greater than in roots (1%) and about nineteen fold greater than the sum of roots and soil (4.1%), a much greater percent recovery than observed in other foliar labeling studies. Averaged over all four foliar split-application rates, ¹⁵N recovery by Desmodium shoots was greater than Pueraria. Results demonstrate that ¹⁵N foliar application to legumes is an effective method for labeling, resulting in atom% excess ¹⁵N levels and ¹⁵N recoveries comparable to those reported with the more traditional soil-labeling approach. Another advantage of this method is a nondestructive, in situ labeling method that permits separation of shoot and root residual N contribution to subsequent crops in N tracer studies.     

Nitrogen mineralization by bacterial-feeding nematodes: verification and measurement

Bacterial feeding nematodes excrete N assimilated in excess of that required for growth. Because metabolic and developmental rates differ among nematode species, we hypothesized that their contribution to N mineralization in soil would differ. Sand-column microcosms amended with an organic substrate, bacteria, and with or without bacterial-feeding nematodes, were leached at 3-d intervals. Cumulative N, as NH ₄ ⁺ or NO ₃ ^- , leached from columns containing nematodes was consistently greater than from columns without nematodes. Maximum N-mineralization rates for populations of rhabditid nematodes, which predominated in field soils early in the summer were at lower temperatures than those for cephalobid nematodes, which predominated later in the summer. For an organic substrate with C-to-N ratio of 11:1, rates of N mineralization among species of different body size were similar, ranging between 0.0012 and 0.0058 g-N nematode^-1 d^-1, mainly as NH ₄ ⁺ . Smaller nematodes mineralized more N per unit of body weight than larger nematodes. We hypothesized that at low C-to-N ratios of the organic substrate, bacterial growth is C-limited and N-immobilization will be minimal; at high C-to-N ratios bacterial growth will be N-limited and there may be rapid immobilization of newly-mineralized N. Consequently, net N mineralization in the presence of nematodes will be lower when the organic substrate has a high C-to-N ratio. In experiments with different nematode species, net mineralization and the nematode contribution to mineralization generally decreased with increasing C-to-N ratio, consistent with the hypothesis; however, there were exceptions.     

Availability of phosphorus and sulfur of insecticide origin by fungi

Thirteen fungal species isolated from soil treated with pesticides were tested for their ability to mineralize and degrade three organophosphate insecticides currently used in Egypt (Cyolan®, Malathion® and Dursban®) in liquid media free from phosporus (P) and sulfur (S). All fungal species grew successfully on the culture media treated with the three used doses of insecticides (10, 50 and 100 ppm active ingredient) but the growth rate varied with the species, the insecticide and the doses. At 10 ppm level, insecticide degradation expressed in term of organic P mineralization (calculated as % of applied P) was the highest with all fungi tested. Organic P mineralization from pesticides was decreased by increasing the dose used to 50 and 100 ppm. The highest amount of P mineralized was observed with Cyolan® followed by Malathion® whilst P mineralization from Dursban® proceeded very slowly. Aspergillus terreus showed the greatest potential to mineralize organic P followed by A. tamarii, A. niger, Trichoderma harzianum and Penicillium brevicompactum whilst the remaining fungi only moderately mineralized the organic P component of the insecticides tested. Organic sulfur mineralization by the used fungal species paralleled, to some extent, organic P mineralization. The extracellular protein content of culture filtrates in the presence of various doses of insecticides was also decreased by increasing insecticide concentrations. The extracellular protein was significantly correlated with P and S mineralization (r = 0.89** and 0.64**, respectively) whilst correlation with cell dry mass was not significant (r = 0.03 and 0.003) suggesting a direct relationship between pesticide degradation and microbial protein production. The addition of P or S to the growth media enhanced extracellular protein excretion, and increased organic P and S mineralization by the most potent species tested (A. niger, A. tamarii, A. terreus and T. harzianum). This increment was significant in most cases, especially at the higher application rates. The relationship between extracellular protein excretion and organic P and S mineralization from insecticides was highly significant with the addition of inorganic phosphorus (r = 0.96** and 0.83**, respectively) or sulfur (r = 0.85** and 0.89**, respectively) to the growth media.     

Fate of explosives and their metabolites in bioslurry treatment processes

Microcosm tests simulating bioslurry reactors with 40% soilcontent, containing high concentrations of TNT and/or RDX,and spiked with either [¹⁴C]-TNT or[¹⁴C]-RDX were conducted to investigate the fate ofexplosives and their metabolites in bioslurry treatment processes.RDX is recalcitrant to indigenous microorganisms in soil andactivated sludge under aerobic conditions. However, soilindigenous microorganisms alonewere able to mineralize 15% of RDX to CO₂ underanaerobic condition, and supplementation of municipal anaerobicsludge as an exogenous source of microorganismssignificantly enhanced the RDX mineralization to 60%. RDXmineralizing activity of microorganisms in soil and sludge wassignificantly inhibited by the presence of TNT. TNTmineralization was poor (< 2%) and was not markedlyimproved by the supplement ofaerobic or anaerobic sludge. Partitioning studies of[¹⁴C]-TNT in the microcosmsrevealed that the removal of TNTduring the bioslurry process was due mainly to thetransformation of TNT and irreversiblebinding of TNT metabolites onto soil matrix. In the case ofRDX under anaerobic conditions,a significant portion (35%) of original radioactivity wasalso incorporated into the biomass andbound to the soil matrix.     

Plant Effects on Spatial and Temporal Patterns of Nitrogen Cycling in Shortgrass Steppe

Because of the water-limited nature and discontinuous plant cover of shortgrass steppe, spatial patterns in ecosystem properties are influenced more by the presence or absence of plants than by plant type. However, plant type may influence temporal patterns of nutrient cycling between plant and soil. Plants having the carbon-3 (C₃) or carbon-4 (C₄) photosynthetic pathway differ in phenology as well as other attributes that affect nitrogen (N) cycling. We estimated net N mineralization rates and traced nitrogen-15 (¹⁵N) additions among plant and soil components during May, July, and September of 1995 in native plots of C₃ plants, C₄ plants, or mixtures of C₃ and C₄. Net N mineralization was significantly greater in C₃ plots than in C₄ plots during both July and September. C₃ plots retained significantly more ¹⁵N in May than did mixed and C₄ plots; these differences in ¹⁵N retention were due to greater ¹⁵N uptake by C₃ plants than by C₄ plants during May. There were no significant differences in total ¹⁵N retention among plant communities for July and September. Soil ¹⁵N was influenced more by presence or absence of plants than by type of plant; greater quantities of ¹⁵N remained in soil interspaces between plants than in soil directly under plants for July and September. Our results indicate that plant functional type (C₃ versus C₄) can affect both the spatial and the temporal patterns of N cycling in shortgrass steppe. Further research is necessary to determine how these intraseasonal differences translate to longer-term and coarser-scale effects of plants on N cycling, retention, and storage. Received 8 December 1997; accepted 6 May 1998.     

Effects of arachidonic acid and docosahexaenoic acid on differentiation and mineralization of MC3T3‐E1 osteoblast‐like cells

Osteoblasts in culture can differentiate into mature mineralizing osteoblasts when stimulated with osteogenic agents. Clinical trials and in vivo animal studies suggest that specific polyunsaturated fatty acids (PUFAs) may benefit bone health. The aim of this study was to investigate whether arachidonic acid (AA) and docosahexaenoic acid (DHA) affect osteogenesis in osteoblasts and the transdifferentiation into adipocytes. Results from this study show that long‐term exposure to AA inhibited alkaline phosphatase (ALP) activity in these cells, which might be prostaglandin E₂ (PGE₂)‐mediated. DHA exposure also inhibited ALP activity which was evident after both short‐ and long‐term exposures. The mechanism whereby DHA inhibits ALP activity is not clear and needs to be investigated. Although long‐term exposure to PUFAs inhibited ALP activity, the mineralizing properties of these cells were not compromised. Furthermore, PUFA exposure did not induce adipocyte‐like features in these cells as evidenced by the lack of cytoplasmic triacylglycerol accummulation. More research is required to elucidate the cellular mechanisms of action of PUFAs on bone. Copyright © 2008 John Wiley & Sons, Ltd.